RESUMO

Pollinators are important drivers of angiosperm diversification at both micro- and macroevolutionary scales. Both hummingbirds and bats pollinate the species-rich and morphologically diverse genus Vriesea across its distribution in the Brazilian Atlantic Forest. Here, we (i) determine if floral traits predict functional groups of pollinators as documented, confirming the pollination syndromes in Vriesea and (ii) test if genetic structure in Vriesea is driven by geography (latitudinal and altitudinal heterogeneity) or ecology (pollination syndromes). We analysed 11 floral traits of 58 Vriesea species and performed a literature survey of Vriesea pollination biology. The genealogy of haplotypes was inferred and phylogenetic analyses were performed using chloroplast (rps16-trnk and matK) and nuclear (PHYC) molecular markers. Floral traits accurately predict functional groups of pollinators in Vriesea. Genetic groupings match the different pollination syndromes. Species with intermediate position were found between the groups, which share haplotypes and differ morphologically from the typical hummingbird- and bat-pollinated flowers of Vriesea. The phylogeny revealed moderately to well-supported clades which may be interpreted as species complexes. Our results suggest a role of pollinators driving ecological isolation in Vriesea clades. Incipient speciation and incomplete lineage sorting may explain the overall low genetic divergence within and among morphologically defined species, precluding the identification of clear species boundaries. The intermediate species with mixed floral types likely represent a window into shifts between pollinator syndromes. This study reports the morphological-genetic continuum that may be typical of ongoing pollinator-driven speciation in biodiversity hotspots.

RESUMO

Network analysis is a powerful tool to understand community-level plant-pollinator interactions. We evaluated the role of floral visitors on plant fitness through a series of pollination exclusion experiments to test the effectiveness of pollinators of an Ipomoea community in the Pacific coast of Mexico, including: (1) all flower visitors, (2) visitors that contact the reproductive organs, (3) visitors that deposit pollen on stigmas, and (4) visitors that mediate fruit and seed production. Our results show that networks built from effective pollination interactions are smaller, less connected, more specialized and modular than floral visitor networks. Modules are associated with pollinator functional groups and they provide strong support for pollination syndromes only when non-effective interactions are excluded. In contrast to other studies, the analyzed networks are not nested. Our results also show that only 59% of floral visitors were legitimate pollinators that contribute to seed production. Furthermore, only 27% of the links in visitation network resulted in seed production. Our study shows that plant-pollination networks that consider effectiveness measures of pollination in addition to floral visitation provide insightful information about the different role floral visitors play in a community, encompassing a large number of commensalistic/antagonistic interactions and the more restricted set of mutualistic relationships that underlie the evolution of convergent floral phenotypes in plants.

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RESUMO

PREMISE: Closely related plant species with overlapping ranges often experience competition for pollination services. Such competition can select for divergence in floral traits that attract pollinators or determine pollen placement. While most species in Centropogon (Campanulaceae: Lobelioideae) have flowers that suggest adaptation to bat or hummingbird pollination, actual pollinators are rarely documented, and a few species have a mix of traits from both pollination syndromes. We studied the pollination biology of a "mixed-syndrome" species and its co-occurring congeners to examine the relationship between floral traits and visitation patterns for Centropogon. METHODS: Fieldwork at two sites in Bolivian cloud forests involved filming floral visitors, quantifying pollen transfer, and measuring floral traits. Stamen exsertion, which determines pollen placement, was measured from herbarium specimens across the geographic range of these species to test for character displacement. RESULTS: Results show a generalization gradient, from primarily bat pollination in white-flowered Centropogon incanus, to bat pollination with secondary hummingbird pollination in the cream-flowered C. brittonianus, to equal reliance on both pollinators in the red-flowered, mixed-syndrome C. mandonis. Pollen transfer between these species is further reduced by differences in stamen exsertion that are accentuated in zones of sympatry, a pattern consistent with character displacement. CONCLUSIONS: Our results demonstrate that key differences in floral color and shape mediate a gradient of specialization in Bolivian Centropogon. Interspecific pollen transfer is further reduced by potential character displacement of a key trait. Broadly, our results have implications for understanding the hyper-diversity of Andean cloud forests, in which multiple species of the same genus frequently co-occur.

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RESUMO

Most angiosperms rely on animal pollination for reproduction, but the dependence on specific pollinator groups varies greatly between species and localities. Notably, such dependence may be influenced by both floral traits and environmental conditions. Despite its importance, their joint contribution has rarely been studied at the assemblage level. At two elevations on the Caribbean island of Dominica, we measured the floral traits and the relative contributions of insects versus hummingbirds as pollinators of plants in the Rubiaceae family. Pollinator importance was measured as visitation rate (VR) and single visit pollen deposition (SVD), which were combined to assess overall pollinator effectiveness (PE). In the wet and cool Dominican highland, we found that hummingbirds were relatively more frequent and effective pollinators than insects, whereas insects and hummingbirds were equally frequent and effective pollinators at the warmer and less rainy midelevation. Furthermore, floral traits correlated independently of environment with the relative importance of pollinators, hummingbirds being more important in plant species having flowers with long and wide corollas producing higher volumes of dilute nectar. Our findings show that both environmental conditions and floral traits influence whether insects or hummingbirds are the most important pollinators of plants in the Rubiaceae family, highlighting the complexity of plant-pollinator systems.

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RESUMO

Flowers show important structural variation as reproductive organs but the evolutionary forces underlying this diversity are still poorly understood. In animal-pollinated species, flower shape is strongly fashioned by selection imposed by pollinators, which is expected to vary according to guilds of effective pollinators. Using the Antillean subtribe Gesneriinae (Gesneriaceae), we tested the hypothesis that pollination specialists pollinated by one functional type of pollinator have maintained more similar corolla shapes through time due to more constant and stronger selection constraints compared to species with more generalist pollination strategies. Using geometric morphometrics and evolutionary models, we showed that the corolla of hummingbird specialists, bat specialists, and species with a mixed-pollination strategy (pollinated by hummingbirds and bats; thus a more generalist strategy) have distinct shapes and that these shapes have evolved under evolutionary constraints. However, we did not find support for greater disparity in corolla shape of more generalist species. This could be because the corolla shape of more generalist species in subtribe Gesneriinae, which has evolved multiple times, is finely adapted to be effectively pollinated by both bats and hummingbirds. These results suggest that ecological generalization is not necessarily associated with relaxed selection constraints.

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RESUMO

The tropical Andes of South America, the world's richest biodiversity hotspot, are home to many rapid radiations. While geological, climatic, and ecological processes collectively explain such radiations, their relative contributions are seldom examined within a single clade. We explore the contribution of these factors by applying a series of diversification models that incorporate mountain building, climate change, and trait evolution to the first dated phylogeny of Andean bellflowers (Campanulaceae: Lobelioideae). Our framework is novel for its direct incorporation of geological data on Andean uplift into a macroevolutionary model. We show that speciation and extinction are differentially influenced by abiotic factors: speciation rates rose concurrently with Andean elevation, while extinction rates decreased during global cooling. Pollination syndrome and fruit type, both biotic traits known to facilitate mutualisms, played an additional role in driving diversification. These abiotic and biotic factors resulted in one of the fastest radiations reported to date: the centropogonids, whose 550 species arose in the last 5 million yr. Our study represents a significant advance in our understanding of plant evolution in Andean cloud forests. It further highlights the power of combining phylogenetic and Earth science models to explore the interplay of geology, climate, and ecology in generating the world's biodiversity.

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RESUMO

Many mistletoe species produce 'bird'-pollinated flowers; however, the reproductive biology of the majority of these species has not been studied. Psittacanthus auriculatus is a Mexican endemic mistletoe, most common in open, dry mesquite grassland. Knowledge of the reproductive biology of P. auriculatus is essential for understanding species formation and diversification of Psittacanthus mistletoes, but it is currently poorly understood. Thus, we studied floral biology and phenology, nectar production and breeding system and pollination of this species. The hermaphroditic red-pink flowers open from the middle to the tip and petals are curly, but remain partially fused forming a floral tube of ca. 20-mm long. Flowers are partially protandrous, produce large amounts of nectar, last 2 days, and stigma receptivity is highest during the second day. We recorded hummingbirds (Cynanthus latirostris, Hylocharis leucotis, Amazilia beryllina, A. violiceps, Calothorax lucifer, Archilochus colubris) and less commonly butterflies (Agraulis vanillae, Anteos clorinde, Papilio multicaudatus, Phocides urania, Phoebis sennae) as floral visitors. P. auriculatus flowers are self-compatible. However, this mistletoe is an obligate animal-pollinated species, as the sensitive stigma avoids self-pollination. Under natural conditions, reproductive success was higher than in manually selfed or cross-pollinated flowers, likely due to the traplining foraging behaviour of hummingbirds. We suggest that the apparent efficient foraging behaviour of hummingbirds maintains gene flow among P. auriculatus, promoting outcrossing.

Assuntos

RESUMO

Pollination syndromes are defined as suites of floral traits evolved in response to selection imposed by a particular group of pollinators (e.g., butterflies, hummingbirds, bats). Although numerous studies demonstrated their occurrence in plants pollinated by radically different pollinators, it is less known whether it is possible to identify them within species pollinated by one functional pollinator group. In such a framework, we expect floral traits to evolve also in response to pollinator subgroups (e.g., species, genera) within that unique functional group. On this, specialised pollination systems represent appropriate case studies to test such expectations. Calceolaria is a highly diversified plant genus pollinated by oil-collecting bees in genera Centris and Chalepogenus. Variation in floral traits in Calceolaria has recently been suggested to reflect adaptations to pollinator types. However, to date no study has explicitly tested that observation. In this paper, we quantitatively test that hypothesis by evaluating the presence of pollination syndromes within the specialised pollination system formed by several Calceolaria and their insect pollinators. To do so, we use multivariate approaches and explore the structural matching between the morphology of 10 Calceolaria taxa and that of their principal pollinators. Our results identify morphological matching between floral traits related to access to the reward and insect traits involved in oil collection, confirming the presence of pollinator syndromes in Calceolaria. From a general perspective, our findings indicate that the pollination syndrome concept can be also extended to the intra-pollinator group level.

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RESUMO

The hummingbird-visited plant community located on the open-habitat mountaintop of the Espinhaço Range was studied for two years (from August 2007 to July 2009) in Serra do Cipó National Park, Southeastern Brazil (19° 15′ S and 43° 31′ W). The floral characteristics and flowering period of the hummingbird-visited plants was monthly recorded along trails located in three vegetation types: (1) typical campos rupestres (TCR), (2) open fields (OPF), and (3) capões de mata (CAM). Hummingbird visitation was observed in 51 plant species, 22 ornithophilous and 29 non-ornithophilous species. The TCR showed the greatest number of species visited (N = 38), followed by the OPF (N = 18) and CAM (N = 17). Six species of hummingbirds were recorded visiting flowers: Augastes scutatus, Campylopterus largipennis, Colibri serrirostris, Chlorostilbon lucidus, Eupetomena macroura and Phaethornis pretrei. This study demonstrates that the species richness and the number of ornithophilous species visited by the hummingbirds at the study site are more similar to hummingbird-plant communities of the Atlantic Forest than to those of the Cerrado communities and other Brazilian highland open-habitat communities. The plant families most visited by hummingbirds were Bromeliaceae and Asteraceae. Although the Asteraceae family is rarely used as a food resource for hummingbirds in other high and lowland communities, in the study site this family is used mainly by the endemic hummingbird Augastes scutatus. We found a large overlap of flowering throughout the year among the species visited by the hummingbirds. Thus, the nectar availability supports these resident hummingbirds. The present study also showed that the studied hummingbird-plant community is composed of many species endemic to the campos rupestres of the Espinhaço Range, some of which are considered to be in danger of extinction, thus constituting a unique and threatened community. Thus, understanding hummingbird-plant pollination dynamics becomes fundamental to the conservation of the campos rupestres.

A comunidade de flores visitadas por beija-flores em habitats abertos de montanhas da Serra do Espinhaço, sudeste do Brasil foi estudada por dois anos (de agosto de 2007 a julho de 2009) no Parque Nacional da Serra do Cipó (19° 15′ S e 43° 31′ W). As características florais e o período de floração das plantas visitadas foram registradas mensalmente ao longo de trilhas localizadas em três tipos vegetacionais: (1) campos rupestres típicos (TCR), (2) campos abertos (OPF), e (3) capões de mata (CAM). Foram observadas visitas de beija-flores a 51 espécies de plantas: 22 ornitófilas e 29 não-ornitófilas. O TCR apresentou o maior número de espécies visitadas (N = 38), seguido pelo OPF (N = 18) e CAM (N = 17). Seis espécies de beija-flores foram observadas visitando as flores: Augastes scutatus, Campylopterus largipennis, Colibri serrirostris, Chlorostilbon lucidus, Eupetomena macroura e Phaethornis pretrei. A riqueza de espécies e o número de espécies ornitófilas visitadas pelos beija-flores, neste estudo, foi mais similar à comunidades de plantas visitadas por beija-flores na Floresta Atlântica, que as comunidades do Cerrado, bem como de outras comunidades de ambientes abertos do Brasil. As famílias com maior número de plantas visitadas pelos beija-flores foram Bromeliaceae e Asteraceae. Apesar da família Asteraceae, raramente ser utilizada como fonte alimentar pelos beija-flores em outras comunidades, nas áreas amostradas neste estudo esta família foi utilizada, principalmente, pelo beija-flor endêmico A. scutatus. Nós registramos grande sobreposição de floração ao longo do ano entre as espécies visitadas pelos beija-flores. Portanto, a disponibilidade de néctar mantem os beija-flores residentes. Este estudo, demonstrou também, que a comunidade de plantas visitadas pelos beija-flores nesta área é composta por muitas espécies endêmicas dos campos rupestres da Serra do Espinhaço, algumas das quais consideradas em perigo de extinção, constituindo assim uma comunidade única e ameaçada. Portanto, o entendimento da dinâmica de polinização desta comunidade é fundamental para a conservação dos campos rupestres.

Assuntos

RESUMO

The hummingbird-visited plant community located on the open-habitat mountaintop of the Espinhaço Range was studied for two years (from August 2007 to July 2009) in Serra do Cipó National Park, Southeastern Brazil (19° 15′ S and 43° 31′ W). The floral characteristics and flowering period of the hummingbird-visited plants was monthly recorded along trails located in three vegetation types: (1) typical campos rupestres (TCR), (2) open fields (OPF), and (3) capões de mata (CAM). Hummingbird visitation was observed in 51 plant species, 22 ornithophilous and 29 non-ornithophilous species. The TCR showed the greatest number of species visited (N = 38), followed by the OPF (N = 18) and CAM (N = 17). Six species of hummingbirds were recorded visiting flowers: Augastes scutatus, Campylopterus largipennis, Colibri serrirostris, Chlorostilbon lucidus, Eupetomena macroura and Phaethornis pretrei. This study demonstrates that the species richness and the number of ornithophilous species visited by the hummingbirds at the study site are more similar to hummingbird-plant communities of the Atlantic Forest than to those of the Cerrado communities and other Brazilian highland open-habitat communities. The plant families most visited by hummingbirds were Bromeliaceae and Asteraceae. Although the Asteraceae family is rarely used as a food resource for hummingbirds in other high and lowland communities, in the study site this family is used mainly by the endemic hummingbird Augastes scutatus. We found a large overlap of flowering throughout the year among the species visited by the hummingbirds. Thus, the nectar availability supports these resident hummingbirds. The present study also showed that the studied hummingbird-plant community is composed of many species endemic to the campos rupestres of the Espinhaço Range, some of which are considered to be in danger of extinction, thus constituting a unique and threatened community. Thus, understanding hummingbird-plant pollination dynamics becomes fundamental to the conservation of the campos rupestres.(AU)

A comunidade de flores visitadas por beija-flores em habitats abertos de montanhas da Serra do Espinhaço, sudeste do Brasil foi estudada por dois anos (de agosto de 2007 a julho de 2009) no Parque Nacional da Serra do Cipó (19° 15′ S e 43° 31′ W). As características florais e o período de floração das plantas visitadas foram registradas mensalmente ao longo de trilhas localizadas em três tipos vegetacionais: (1) campos rupestres típicos (TCR), (2) campos abertos (OPF), e (3) capões de mata (CAM). Foram observadas visitas de beija-flores a 51 espécies de plantas: 22 ornitófilas e 29 não-ornitófilas. O TCR apresentou o maior número de espécies visitadas (N = 38), seguido pelo OPF (N = 18) e CAM (N = 17). Seis espécies de beija-flores foram observadas visitando as flores: Augastes scutatus, Campylopterus largipennis, Colibri serrirostris, Chlorostilbon lucidus, Eupetomena macroura e Phaethornis pretrei. A riqueza de espécies e o número de espécies ornitófilas visitadas pelos beija-flores, neste estudo, foi mais similar à comunidades de plantas visitadas por beija-flores na Floresta Atlântica, que as comunidades do Cerrado, bem como de outras comunidades de ambientes abertos do Brasil. As famílias com maior número de plantas visitadas pelos beija-flores foram Bromeliaceae e Asteraceae. Apesar da família Asteraceae, raramente ser utilizada como fonte alimentar pelos beija-flores em outras comunidades, nas áreas amostradas neste estudo esta família foi utilizada, principalmente, pelo beija-flor endêmico A. scutatus. Nós registramos grande sobreposição de floração ao longo do ano entre as espécies visitadas pelos beija-flores. Portanto, a disponibilidade de néctar mantem os beija-flores residentes. Este estudo, demonstrou também, que a comunidade de plantas visitadas pelos beija-flores nesta área é composta por muitas espécies endêmicas dos campos rupestres da Serra do Espinhaço, algumas das quais consideradas em perigo de extinção, constituindo assim uma comunidade única e ameaçada. Portanto, o entendimento da dinâmica de polinização desta comunidade é fundamental para a conservação dos campos rupestres.(AU)

Assuntos

RESUMO

To describe plant phenological patterns and correlate functioning for the quantity and quality of resources available for the pollinator, it is crucial to understand the temporal dynamics of biological communities. In this way, the pollination syndromes of 46 species with different growth habits (trees, shrubs, herbs, and vines) were examined in an area of Caatinga vegetation, northeastern Brazil (7° 28′ 45″ S and 36° 54′ 18″ W), during two years. Flowering was monitored monthly in all the species, over two years (from January 2003 to December 2004). Pollination syndromes were characterised based on floral traits such as size, colour, morphology, symmetry, floral resources, as well as on direct visual observation of floral visitors on focal plants and published information. We observed differences among the plant growth habits with respect to floral traits, types of resources offered, and floral syndromes. The flowering periods of the species varied among floral syndrome groups. The majority of the melittophilous species flowered during the rainy season in the two study years, while the species of the other pollination syndroms flowered at the end of the dry season. An asynchrony of flowering was noted among the chiropterophilous species, while the phalenophilous group concentrated during the rainy season. The overall availability of floral resources was different during the rainy and the dry seasons, and also it varied among plants with different growth habits. The availability of oil-flowers coincided with the period of low nectar availability. We observed a relationship between the temporal distribution of the pollination syndromes and the availability of floral resources among each growth habits in this tropical ecosystem. Resource allocation in seasonal environments, such as the Caatinga, can function as a strategy for maintaining pollinators, facilitating therefore the reproductive success of plant species. The availability of floral resources during all the year, specially in seasonal environments such as the Caatinga, may function as a strategy to maintain pollinator populations ensuring the reproductive success of the plants.

Descrever o padrão fenológico das plantas e correlacionar com a quantidade e qualidade dos recursos disponíveis para os polinizadores é fundamental para entender a dinâmica temporal das comunidades biológicas. Neste sentido, foram estudadas as síndromes de polinização de 46 espécies com diferentes hábitos (árvores, arbustos, ervas, trepadeiras) em uma área de caatinga, no Cariri Paraibano no Nordeste do Brasil (7° 28′ 45″ S e 36° 54′ 18″ W) durante dois anos. Para as diferentes espécies foi acompanhado o período de floração, sendo destacada a fase de início e o pico. As síndromes de polinização foram caracterizadas com base nos atributos florais, como tamanho, cor, morfologia, simetria, tipo de recurso, bem como a partir de observações visuais diretas dos visitantes florais em plantas focais e informações de literatura. Foram encontradas diferenças entre os hábitos, relacionadas aos distintos atributos florais, tipo de recurso e síndrome floral. O período de floração das espécies mostrou-se distinto entre os diferentes tipos de síndromes. A maioria das espécies melitófilas floresceu na estação úmida, enquanto as demais no final da estação seca, nos dois anos de estudo. Foi observada assincronia na floração das espécies quiropterófilas e concentração entre as esfingófilas na estação úmida. A disponibilidade de recursos florais apresentou diferenças entre as estações seca e chuvosa, diferindo também entre os hábitos. A oferta de flores de óleo coincidiu com o período de menor disponibilidade de néctar. Foi observada relação entre a distribuição temporal das diferentes síndromes de polinização, juntamente com a disponibilidade dos recursos florais, nos diferentes hábitos para este ecossistema tropical. A alocação de recursos em ambientes sazonais, como a Caatinga estudada, pode funcionar como uma estratégia para manutenção de polinizadores, facilitando, portanto o sucesso reprodutivo das espécies vegetais.

Assuntos

RESUMO

To describe plant phenological patterns and correlate functioning for the quantity and quality of resources available for the pollinator, it is crucial to understand the temporal dynamics of biological communities. In this way, the pollination syndromes of 46 species with different growth habits (trees, shrubs, herbs, and vines) were examined in an area of Caatinga vegetation, northeastern Brazil (7° 28′ 45″ S and 36° 54′ 18″ W), during two years. Flowering was monitored monthly in all the species, over two years (from January 2003 to December 2004). Pollination syndromes were characterised based on floral traits such as size, colour, morphology, symmetry, floral resources, as well as on direct visual observation of floral visitors on focal plants and published information. We observed differences among the plant growth habits with respect to floral traits, types of resources offered, and floral syndromes. The flowering periods of the species varied among floral syndrome groups. The majority of the melittophilous species flowered during the rainy season in the two study years, while the species of the other pollination syndroms flowered at the end of the dry season. An asynchrony of flowering was noted among the chiropterophilous species, while the phalenophilous group concentrated during the rainy season. The overall availability of floral resources was different during the rainy and the dry seasons, and also it varied among plants with different growth habits. The availability of oil-flowers coincided with the period of low nectar availability. We observed a relationship between the temporal distribution of the pollination syndromes and the availability of floral resources among each growth habits in this tropical ecosystem. Resource allocation in seasonal environments, such as the Caatinga, can function as a strategy for maintaining pollinators, facilitating therefore the reproductive success of plant species. The availability of floral resources during all the year, specially in seasonal environments such as the Caatinga, may function as a strategy to maintain pollinator populations ensuring the reproductive success of the plants.(AU)

Descrever o padrão fenológico das plantas e correlacionar com a quantidade e qualidade dos recursos disponíveis para os polinizadores é fundamental para entender a dinâmica temporal das comunidades biológicas. Neste sentido, foram estudadas as síndromes de polinização de 46 espécies com diferentes hábitos (árvores, arbustos, ervas, trepadeiras) em uma área de caatinga, no Cariri Paraibano no Nordeste do Brasil (7° 28′ 45″ S e 36° 54′ 18″ W) durante dois anos. Para as diferentes espécies foi acompanhado o período de floração, sendo destacada a fase de início e o pico. As síndromes de polinização foram caracterizadas com base nos atributos florais, como tamanho, cor, morfologia, simetria, tipo de recurso, bem como a partir de observações visuais diretas dos visitantes florais em plantas focais e informações de literatura. Foram encontradas diferenças entre os hábitos, relacionadas aos distintos atributos florais, tipo de recurso e síndrome floral. O período de floração das espécies mostrou-se distinto entre os diferentes tipos de síndromes. A maioria das espécies melitófilas floresceu na estação úmida, enquanto as demais no final da estação seca, nos dois anos de estudo. Foi observada assincronia na floração das espécies quiropterófilas e concentração entre as esfingófilas na estação úmida. A disponibilidade de recursos florais apresentou diferenças entre as estações seca e chuvosa, diferindo também entre os hábitos. A oferta de flores de óleo coincidiu com o período de menor disponibilidade de néctar. Foi observada relação entre a distribuição temporal das diferentes síndromes de polinização, juntamente com a disponibilidade dos recursos florais, nos diferentes hábitos para este ecossistema tropical. A alocação de recursos em ambientes sazonais, como a Caatinga estudada, pode funcionar como uma estratégia para manutenção de polinizadores, facilitando, portanto o sucesso reprodutivo das espécies vegetais.(AU)

Assuntos